Ci a ion: Ma ías, J.; Rod íguez, M.J.;
Ca illo-Vico, A.; Casals, J.; Fonde illa,
S.; Ha os, C.M.; Ped oche, J.; Apa icio,
N.; Fe nández-Ga cía, N.;
Aguiló-Aguayo, I.; e al. F om ‘Fa m
o Fo k’: Explo ing he Po en ial o
Nu ien -Rich and S ess-Resilien
Eme gen C ops o Sus ainable and
Heal hy Food in he Medi e anean
Region in he Face o Clima e Change
Challenges. Plan s 2024,13, 1914.
h ps://doi.o g/10.3390/
plan s13141914
Academic Edi o : Osca Vicen e
Recei ed: 10 Ma ch 2024
Re ised: 8 Ap il 2024
Accep ed: 8 July 2024
Published: 11 July 2024
Copy igh : © 2024 by he au ho s.
Licensee MDPI, Basel, Swi ze land.
This a icle is an open access a icle
dis ibu ed unde he e ms and
condi ions o he C ea i e Commons
A ibu ion (CC BY) license (h ps://
c ea i ecommons.o g/licenses/by/
4.0/).
plan s
Re iew
F om ‘Fa m o Fo k’: Explo ing he Po en ial o Nu ien -Rich
and S ess-Resilien Eme gen C ops o Sus ainable and Heal hy
Food in he Medi e anean Region in he Face o Clima e
Change Challenges
Ja ie Ma ías 1, Ma ía JoséRod íguez 2, An onio Ca illo-Vico 3,4 , Joan Casals 5, Sa a Fonde illa 6,
Claudia Mónika Ha os 7, Jus o Ped oche 8, Nie es Apa icio 9, Nie es Fe nández-Ga cía10,
Ing id Aguiló-Aguayo 11, C is ina Sole -Ri as 12,13, Ped o A. Caballe o 14 , Asunción Mo e 15 , Daniel Rico 16
and Ma ía Regue a 17,*,† on behal o he Nu iC op Spanish Conso ium/G oup
1Ag a ian Resea ch Ins i u e “La O den-Valdeseque a” o Ex emadu a (CICYTEX),
06187 Guadaji a (Badajoz), Spain; ja ie [email p o ec ed]
2Technological Ins i u e o Food and Ag icul u e o Ex emadu a (INTAEX-CICYTEX), A da. Adol o Suá ez
s/n, 06007 Badajoz, Spain; [email p o ec ed]
3Ins i u o de Biomedicina de Se illa, IBiS/Hospi al Uni e si a io Vi gen del Rocío/CSIC/Uni e sidad de
Se illa, 41013 Se ille, Spain; [email p o ec ed]
4Depa amen o de Bioquímica Médica y Biología Molecula e Inmunología, Facul ad de Medicina,
Uni e sidad de Se illa, 41009 Se ille, Spain
5FundacióMiquel Agus í/Ho PTA, Depa men o Ag i-Food Enginee ing and Bio echnology, Uni e si a
Poli ècnica de Ca alunya (UPC)-Ba celonaTech, 08860 Cas ellde els, Spain; [email p o ec ed]
6
Ins i u e o Sus ainable Ag icul u e, Consejo Supe io de In es igaciones Cien í icas, A da. Menéndez Pidal
s/n, 14004 Có doba, Spain; [email p o ec ed]
7Ce eal G oup, Ins i u e o Ag ochemis y and Food Technology (IATA-CSIC), A . Agus ín Esca dino 7,
Pa que Cien í ico, 46980 Valencia, Spain; cmha [email p o ec ed]
8G oup o Plan P o eins, Ins i u o de la G asa, CSIC. C a. de U e a Km. 1, 41013 Se ille, Spain;
[email p o ec ed]
9Ag o-Technological Ins i u e o Cas illa y León (ITACyL), C a. Bu gos Km. 119, 47071 Valladolid, Spain;
[email p o ec ed]
10 Depa men o Abio ic S ess and Plan Pa hology, Cen o de Eda ología y Biología Aplicada del
Segu a (CSIC), Campus Uni e si a io de Espina do, 30100 Mu cia, Spain; [email p o ec ed]
11 Pos ha es P og amme, Ins i u e o Ag i ood Resea ch and Technology (IRTA), Pa c Ag obio ech Lleida,
Pa c de Ga deny, Edi ici F ui cen e, 25003 Lleida, Spain; [email p o ec ed]
12 Depa amen o de P oducción y Ca ac e ización de Nue os Alimen os, Ins i u e o Food Science
Resea ch-CIAL (UAM+CSIC), Campus de Can oblanco, Uni e sidad Au ónoma de Mad id,
C/Nicolas Cab e a 9, 28049 Mad id, Spain; [email p o ec ed]
13 Sección Depa amen al de Ciencias de la Alimen ación, Facul ad de Ciencias,
Uni e sidad Au ónoma de Mad id, Campus de Can oblanco, 28049 Mad id, Spain
14 Food Technology, Depa men o Ag icul u e and Fo es y Enginee ing, Uni e sidad de Valladolid,
34004 Palencia, Spain; ped oan onio.caballe [email p o ec ed]
15 Depa amen o Biología Vege al, Facul ad de Biología, Campus Uni e si a io de Espina do,
Uni e sidad de Mu cia, 30100 Mu cia, Spain; [email p o ec ed]
16 Depa men o Medicine, De ma ology and Toxicology, Uni e sidad de Valladolid, A . Ramón y Cajal, 7,
47005 Valladolid, Spain; [email p o ec ed]
17 Depa amen o de Biología, Campus de Can oblanco, Uni e sidad Au ónoma de Mad id, C/Da win 2,
28049 Mad id, Spain
*Co espondence: ma ia. [email p o ec ed]; Tel.: +34-91497-81-89
†Collabo a o s o he Nu iC op Spanish Conso ium/G oup a e p o ided in he Acknowledgmen s.
Abs ac : In he dynamic landscape o ag icul u e and ood science, inco po a ing eme gen c ops
appea s as a pionee ing solu ion o di e si ying ag icul u e, unlocking possibili ies o sus ainable
cul i a ion and nu i ional bols e ing ood secu i y, and c ea ing economic p ospec s amid e ol ing
en i onmen al and ma ke condi ions wi h posi i e impac s on human heal h. This e iew explo es
he po en ial o u ilizing eme gen c ops in Medi e anean en i onmen s unde cu en clima e
scena ios, emphasizing he mani old bene i s o ag icul u al and ood sys em di e si ica ion and
Plan s 2024,13, 1914. h ps://doi.o g/10.3390/plan s13141914 h ps://www.mdpi.com/jou nal/plan s
Plan s 2024,13, 1914 2 o 45
assessing he impac o en i onmen al ac o s on hei quali y and consume heal h. Th ough a deep
explo a ion o he esilience, nu i ional alue, and heal h impac s o neglec ed and unde u ilized
species (NUS) such as quinoa, ama an h, chia, mo inga, buckwhea , mille , e , hemp, o dese
u les, hei capaci y o h i e in he changing Medi e anean clima e is highligh ed, o e ing no el
oppo uni ies o ag icul u e and unc ional ood de elopmen . By analysing how p omo ing ag i-
cul u al di e si ica ion can enhance ood sys em adap abili y o e ol ing en i onmen al condi ions,
os e ing sus ainabili y and esilience, we discuss ecen indings ha unde sco e he main bene i s
and limi a ions o hese c ops om ag icul u al, ood science, and heal h pe spec i es, all c ucial
o esponsible and sus ainable adop ion. Thus, by using a sus ainable and holis ic app oach, his
e ision analyses how he in eg a ion o NUS c ops in o Medi e anean ag i ood sys ems can enhance
ag icul u e esilience and ood quali y add essing en i onmen al, nu i ional, biomedical, economic,
and cul u al dimensions, he eby mi iga ing he isks associa ed wi h monocul u e p ac ices and
bols e ing local economies and li elihoods unde new clima e scena ios.
Keywo ds: NUS c ops; Medi e anean ag i ood sys ems; abio ic s esso s; ag icul u al di e si ica ion;
nu i ional quali y; heal hy die s; clima e esilience
1. In oduc ion
Conside ing he cu en ends in popula ion g ow h, achie ing global ood secu i y
emains a u u e challenge in which ag icul u e plays a key ole [
1
]. Cu en ly, ag icul u e is
based on he monocul u e o a limi ed numbe o species [
2
]. This p ac ice leads o inc eased
agili y o he ag icul u al sys em (e.g., in he ace o bio ic and abio ic s esses), esul ing
in a deple ion o a ailable ood ing edien s, as ou die elies on a ew plan species [
3
,
4
].
Fu he mo e, he e is subs an ial scien i ic e idence indica ing ha mo e di e se ag i ood
sys ems a e gene ally sa e om bo h nu i ional and ag onomical pe spec i es, showing
g ea e esilience o en i onmen al changes and o e ing a b oade a ay o nu i ionally
e ec i e ood p oduc s [
3
]. Indeed, g owing a a ie y o c ops enhances he nu i ional
di e si y o he local die . This is pa icula ly impo an o ce ain communi ies ha ely
hea ily on a ew s aple c ops, helping add ess nu i ional de iciencies and imp o e o e all
heal h [
4
,
5
]. In oducing no el c ops will also o e a me s supplemen a y income s eams,
diminishing dependence on a singula c op and alle ia ing he inancial isks associa ed
wi h ma ke luc ua ions o ad e se wea he condi ions [
6
]. Pa icula ly, ce ain c ops may
be mo e esilien o d ough , hea , o o he clima e- ela ed challenges, p o iding a bu e
agains he impac s o clima e change [
7
]. Also, di e si ying c ops in o a o a ion sys em
helps b eak pes , weed in es a ion, and disease cycles, imp o es soil e ili y, and educes
he isk o soil deg ada ion [
8
] suppo ing sus ainable ag icul u al p ac ices. Likewise, he
di e si ica ion o ou ag icul u al sys em can con ibu e o minimizing ood dependence
ha may be subjec ed, as we ha e expe ienced in ecen imes, o changes linked o
in e na ional con lic s [9].
Cu en ly, wo- hi ds o he plan -based oods we consume p ima ily come om h ee
ce eals, whea , ice, and maize, p o iding 60% o die a y ene gy [
10
]. While he nu i ional
alue o hese c ops is undeniable, hey a e de icien in essen ial nu ien s c ucial o
ensu ing a heal hy and balanced die , such as mine als, p o eins, essen ial amino acids, o
i amins, among o he s [
4
,
11
]. The e o e, he di e si ica ion o ou ood c ops ( o human
and animal consump ion), by including c ops wi h high nu i ional alue, is essen ial o
achie ing a balanced nu ien in ake and p omo ing g ea e sus ainabili y and mode a ion
in he consump ion o animal-o igin oods [12].
The dis inc clima ic pa e ns o he Medi e anean egion a e cha ac e ized by mild,
we win e s and wa m, d y summe s [
13
]. The unique clima ic condi ions and di e se
ecosys ems o hese a eas ace inc easing challenges de i ed om clima e change which a e
linked o he escala ing occu ence o d ough s and ele a ed empe a u es, which di ec ly
impac ag icul u e [
14
]. Also, pes s and diseases may p oli e a e unde al e ed clima e
Plan s 2024,13, 1914 3 o 45
condi ions, posing addi ional challenges o a me s [
15
]. Fu he mo e, ain ed ag icul-
u e, which plays a pi o al ole in sou he n and eas e n Medi e anean coun ies, makes a
subs an ial con ibu ion o ood secu i y in he egion [
16
]. The e o e, he Medi e anean
ag i ood sys em will equi e he de elopmen o adap a ion s a egies o cope wi h he
a o emen ioned clima ic changing condi ions, gi ing special conside a ion o ain ed ag i-
cul u e, pa icula ly in ligh o i s impo ance in ela ion o i s ex en in e ms o su ace a ea.
This may in ol e he cul i a ion o mo e esilien c op species, which include neglec ed
and unde u ilized species (NUS) and imp o ed (b ed) c op a ie ies, changes in a ming
p ac ices, and he implemen a ion o sus ainable ag icul u al echniques o mi iga e he
impac o clima e change, among o he s [10,17].
NUS, a no el and e ol ing ca ego y in ag icul u e, holds p o ound signi icance o
global ood secu i y, pa icula ly in add essing nu i ional challenges. These c ops ep esen
a di e se a ay o plan species ha exhibi esilience and adap abili y. Thei eme gence is a
esponse o he p essing need o sus ainable and nu i ious ood sou ces, mos ly when con-
side ing he challenges aced by con en ional c ops, such as suscep ibili y o pes s, diseases,
and clima e change, especially in suscep ible egions such as he Medi e anean [
13
–
15
].
They a e p omising al e na i es o cul i a ion in di e se ecosys ems and hei adap abili y
can con ibu e o inc eased Medi e anean ag icul u al p oduc i i y [18].
O e all, i should be conside ed ha he widesp ead adop ion o ag icul u al di-
e si ica ion p ac ices in he Medi e anean egion can con ibu e o es ablishing a mo e
sus ainable and e ec i e ag icul u e and ood sys em a bo h local and global scales [
19
].
The cul i a ion o new c ops can boos u al economies by c ea ing employmen , suppo -
ing local businesses, and con ibu ing o o e all economic g ow h. Howe e , i is impo an
o no e ha he success ul in oduc ion o new c ops equi es ca e ul conside a ion o
ac o s such as soil sui abili y, clima e condi ions, ma ke demand, and a me s’ capaci y o
adap o new cul i a ion p ac ices, among o he s [
17
]. Addi ionally, suppo i e policies,
ex ension se ices, and esea ch ini ia i es play a i al ole in acili a ing he success ul
in eg a ion o new c ops [20].
Taking all hese aspec s in o conside a ion, his e iew aims o explo e he po en ial o
nu ien - ich and s ess- esilien eme gen c ops as a iable solu ion o os e sus ainable
and heal hy ood p oduc ion in he ace o clima ic challenges in he Medi e anean egion.
By examining he adap abili y and nu i ional alue o selec ed NUSs (including quinoa
(Chenopodium quinoa Willd.), ama an h (Ama an hus spp.), chia (Sal ia hispanica L.), mo inga
(Mo inga olei e a Lam.), buckwhea (Fagopy um esculen um Moench and Fagopy um a a icum
(L.) Gae n.), mille (including Panicum miliaceum L. o Se a ia i alica L., among o he s), e
(E ag os is e (Zucc.) T o e ), hemp (Cannabis sa i a L.), o dese u les (including Te ezia
cla e yi Cha in)), we aim o shed ligh on hei ole in mi iga ing he ad e se e ec s o
clima e change on ag icul u al sys ems. This wo k no only del es in o he ag onomical
ea u es o hese c ops bu also conside s hei nu i ional, ood echnological, and heal h
cha ac e is ics, emphasizing he impo ance o di e si ying c ops in he Medi e anean
egion o esilien and nu i ionally ich ood sou ces. Th ough his explo a ion, we aspi e
o con ibu e o he de elopmen o s a egies ha p omo e ood secu i y and sus ainabili y
in he Medi e anean egion amids he dynamic landscape o clima e change.
2. Ag onomical Aspec s Rela ed o NUSs o he Medi e anean Region
2.1. His o ical Backg ound
Since he ad en o ag icul u e, he Medi e anean egion has been a cen e o ag icul-
u al inno a ion and c op di e si ica ion [
21
]. Pedoclima ic condi ions, cha ac e ized by a
summe d y season and ligh soils, acili a ed he in oduc ion o Neoli hic c op cul u es
cen ed a ound annual plan s [22]. Ea ly a me s inhe i ed ag icul u al p ac ices om he
Nea Eas , inco po a ing domes ica ed c ops such as whea and ba ley (Ho deum ulga e L.),
along wi h legumes such as peas (Pisum sa i um L.), len ils (Lens culina is L.), and chickpeas
(Cice a ie inum L.) [
23
]. This unde sco es he signi icance o hese ini ial in oduc ions o e
millennia. The p olonged his o y o co-e olu ion be ween hese in oduced plan s, local
Plan s 2024,13, 1914 4 o 45
ag oclima ic condi ions, and human p e e ences has shaped he Medi e anean a ea in o
an impo an seconda y di e si y cen e [24].
Buil on his ounda ion, h oughou his o y, new plan species we e in oduced, being
c ucial o colonizing new c oplands and enhancing ood secu i y. These can be conside ed
he eme ging c ops o hei espec i e e as. Fo ins ance, in ea ly imes, ye (Secale ce eale
L.), ini ially in oduced as a weed, became a signi ican c op o expanding ag icul u al
socie ies in no he n Eu ope due o i s compe i i e s eng h in poo soils and un a ou able
clima es [
25
]. In he Middle Ages, A abic se lemen s in oduced c ops like ice, playing
a i al ole in de eloping ag icul u e in del aic a eas, and so ghum (So ghum bicolo L.
Moench), cul i a ed in ma ginal lands wi h insu icien ain all, con ibu ing o ensu ing
ood p oduc ion [
26
]. In mo e ecen imes, exchanges wi h he Ame icas in oduced key
species, no ably maize and po a o (Solanum ube osum L.), swi ly in eg a ed in o ag icul-
u al sys ems, especially in egions acing challenges in p oducing b ead-making ce eals.
Se e al o he ‘eme ging c ops’ we e success ul in speci ic a eas, highligh ing he
signi icance o plan in oduc ions o local ag icul u e in he pas . Many o hese a e now
conside ed NUS bu hold po en ial as al e na i e c ops o enhance ag icul u al p oduc ion
in speci ic en i onmen s [
27
]. Some o he examples a e common mille (Panicum miliaceum
L.), ox ail mille (Se a ia i alica L.), buckwhea , o di e en T i icum species such as einko n
(T.monococcum L.), polish whea (T.polonicum L.), club whea (T.compac um L.), o spel
(T.spel a L.) [
28
]. These plan s played pi o al oles in ood secu i y du ing pe iods o
ag icul u al c isis, caused by he deple ion o soil e ili y o clima e c isis. A no able
example o he ole o eme ging c ops in adap ing o clima e changes is demons a ed by
socie ies’ esponse o he Li le Ice Age (LIA) cooling (1550–1710) [
29
]. This cooling pe iod
had ad e se e ec s on ag icul u al p oduc ion, leading o heigh ened amine. Socie ies ha
di e si ied ag icul u e, h ough he in oduc ion o new c ops and adap ing cul i a ion
echniques, exhibi ed inc eased esilience [30].
Apa om plan species, ungi ha e also been impo an o human nu i ion. Fo
example, ungal species ha e been ha es ed om he wild o cul i a ed o complemen
die s [
31
]. A no able example is he dese u le (Te ezia cla e yi Cha in), which has ecen ly
been in eg a ed in o Medi e anean c opping sys ems. His o ically, dese u les we e
ga he ed om he wild by di e se cul u es ac oss he Middle Eas , A ica, and Aus alia.
Despi e geog aphical di e ences, hese cul u es used simila me hods o u le collec ion,
p epa a ion, and use [
32
]. The cul i a ion o dese u les began ela i ely ecen ly, wi h
he i s plan a ion es ablished in Mu cia, Spain, in 1999. This e o led o he p oduc ion
o u les wo yea s la e , ma king he s a o global dese u le cul i a ion, which has
since expanded o a ious egions in Spain and o he coun ies [33,34].
In summa y, om he o igin o Medi e anean ag icul u al socie ies o he p esen ,
eme ging c ops ha e been undamen al o ag icul u al de elopmen and ood secu i y.
Adap ing ag icul u e o he cu en clima e c isis should d aw om pas success ul expe-
iences, le e aging he ag onomic po en ial o unde u ilized c ops and he di e si y o
land aces ha ha e co-e ol ed wi h Medi e anean ag osys ems.
2.2. Impo ance and Bene i s
Biodi e si y holds pa amoun impo ance o ensu ing ood secu i y and nu i ion
in Medi e anean ag icul u al ecosys ems. The ich di e si y wi hin ag icul u al sys ems
se es as a ounda ional asse o con inually enhancing a ie ies essen ial o sus aining
ag icul u e in he con ex o clima e change. In line wi h his, he c ucial ole o NUS in
sus ainable ag icul u e, ensu ing imp o ed en i onmen al p o ec ion and a consis en ood
supply, en ails selec ing cos -e ec i e a ie ies adap ed o cu en cul i a ion a eas and o
economic and ag onomic in e es [
1
]. Despi e NUS encompassing c ops ha lack b oad
adop ion, hey o e added alue compa ed o con en ional c ops, being nu ien ich,
adap able o poo soils and a id a eas, equi ing minimal wa e o p ope de elopmen ,
and u ilizing nea ly 100% o he plan ( oo , lea , seed, e c.) o u le.
Plan s 2024,13, 1914 5 o 45
In he ace o clima e change and e ol ing ma ke s, knowledge and inno a ion be-
come pi o al o sus ainable success in ag icul u e [
35
]. Enhancing inno a ion pe o mance
becomes c ucial o ins i u ions and companies in he ag icul u al sec o o emain compe i-
i e. Beyond echnology adop ion, inno a ion mus an icipa e ma ke needs and iden i y
supe io p oduc s and p ocesses, as exempli ied by he in oduc ion o new sus ainable
c ops in Medi e anean a eas such as Po ugal, Spain, I aly, G eece, Tu key, Egyp , and
Mo occo [36].
Recognizing he signi ican nega i e impac o clima e change on ood p oduc ion,
he in e na ional scien i ic communi y acknowledges inc eased isks such as empe a u e
luc ua ions, d ough s, and loods. The ag icul u al sec o , a ne emi e o g eenhouse
gases, aces ulne abili y o clima e change e ec s. Adap ing, educing ulne abili ies,
and inc easing esilience o clima e change is hus c ucial. The in oduc ion o NUS c ops
eme ges as a s a egy o adap and ensu e ood secu i y and good nu i ion in he ace o
clima e change, aligning wi h he 17 Sus ainable De elopmen Goals commi ed by he
global communi y [
37
]. Howe e , clima e change poses challenges o achie ing hese goals,
hinde ing li elihood de elopmen and ood secu i y, especially o ulne able popula ions.
The inco po a ion o NUS becomes ins umen al in o e coming hese challenges and
os e ing ood secu i y and nu i ion o humani y in he u u e [37].
2.3. Classi ica ion and Cha ac e is ics
NUSs a e also e e ed o as no el o eme gen c ops, b eak c ops, o al e na i e
c ops, among o he e ms in use. These c ops can be classi ied based on a ious c i e ia
such as use, clima e adap abili y, nu i ional con en , g ow h equi emen s, and o he
speci ic ai s. Acco ding o hei in ended use, eme ging c ops can be des ined o ood,
e.g., quinoa, o o indus ial use, e.g., hemp. The classi ica ion can also ocus on ai s
associa ed wi h clima e adap abili y, which make c ops well sui ed o speci ic condi ions
(Table 1). Consequen ly, di e en ca ego ies can be es ablished in his line, such as d ough -
ole an c ops (e.g., quinoa [
38
], mille [
39
], hemp [
40
], and dese u les [
41
]), hea -
esis an c ops (e.g., mille [
42
], ama an h [
43
]), o salini y- ole an c ops (e.g., quinoa [
44
],
so ghum [
45
]). Fu he mo e, c op g ow h equi emen s can be ca ego ized in o h ee
le els: low, medium, and high. In he case o NUS, he e is signi ican impo ance placed on
cul i a ing c ops wi h ela i ely low g ow h equi emen s. This emphasis is gea ed owa ds
educing he en i onmen al impac o ag icul u e on clima e change and enhancing he
esilience o ag icul u al sys ems. Mo eo e , eme ging c ops can also be classi ied based
on hei nu i ional alue (See Sec ion 3), es ablishing di e en ca ego ies such as high-
quali y p o ein c ops (e.g., quinoa, ama an h, mo inga, dese u les), nu ien -dense
c ops (e.g., mo inga, quinoa, ama an h), o omega-3- ich c ops (e.g., chia, laxseeds (Linum
usi a issimum L.)), among o he s.
Quinoa, ama an h, mille , buckwhea , hemp, e , mo inga, dese u les, and chia,
among o he s (Figu e 1), a e conside ed unde u ilized clima e- esilien c ops [
46
–
53
]
(
Table 1
). The in oduc ion o hese eco- iendly c ops in o ou ag icul u e has he po en-
ial o enhance biodi e si y and esilience in ag icul u al sys ems. Cu en ly, ag icul u al
p oduc ion hea ily elies on only a ew species, and di e si ying wi h hese c ops can
con ibu e o a mo e sus ainable and obus ag icul u al ecosys em [
51
]. These eme ging
c ops showcase ag onomic ai s well sui ed o di e se en i onmen al condi ions, cha -
ac e is ic o each species. Fo ins ance, quinoa shows g ea po en ial due o i s abili y
o wi hs and abio ic s esses g owing in ha sh en i onmen s wi h low e ili y [
54
]. I s
adap abili y o di e se en i onmen al condi ions is a ibu ed o i s ex ensi e gene ic di-
e si y [
55
], exhibi ing ole ance o d ough , os , and salini y, while h i ing on poo
soils [
54
]. Ama an h, mille , hemp, chia, and dese u le also demons a ed esilience o
d ough [
39
,
40
,
56
–
58
], making hem low-wa e - equi emen op ions o cul i a ion. Te
is a wa m-season ce eal c op adap ed o a wide ange o en i onmen s ha can be g own
unde ain ed condi ions [
59
]. Mo inga exhibi s ole ance o d ough condi ions. In egions

Plan s 2024,13, 1914 6 o 45
wi h a sub opical clima e, mo inga can esis up o six mon hs o d y seasons, p o ided
he annual ain all amoun s o 500 mm [60].
Plan s 2024, 13, x FOR PEER REVIEW 7 o 46
Figu e 1. NUS c ops wi h he po en ial o cul i a ion in Medi e anean en i onmen s. Rep esen a-
i e images o diffe en nu ien - ich and s ess- esilien eme gen c ops o sus ainable and heal hy
ood in he Medi e anean egion, g own unde ield condi ions ac oss diffe en egions o Spain.
The images depic he c ops in he ield (ou e images) and he panicle, in lo escence, o uffle o
each c op (inne image). The c ops included in he igu e a e Chenopodium quinoa Willd., Ama an hus
sp., Fagopy um sp., E ag os is e (Zucc.) T o e , Sal ia hispanica L., Mo inga olei e a Lam., Cannabis sa i a
L., and mille (Pennise um glaucum (L.) R.B . Te ezia cla e yi Cha in (wi h Helian hemum alme iense Pau)).
2.4. Challenges and Limi a ions
The cul i a ion o NUSs poses a ious challenges ac oss diffe en ace s o he ag i-
cul u al and supply chain landscape. Ongoing h ea s o ag icul u al en i onmen s s em
om a g owing wo ld popula ion, limi ed a able land, and swi clima e changes. P ojec-
ions sugges ha he global popula ion will su pass 9 billion by 2050 [73]. The challenge
o eeding his expanding popula ion is a global conce n. To secu e ood and ecosys em
s abili y, i is essen ial o de elop NUSs as c ops o sus ainable ag icul u e, emphasizing
inc eased ne p oduc ion while minimizing ad e se en i onmen al impac s. Ne e he-
less, inco po a ing NUSs in o exis ing o a ions in oduces complexi ies, necessi a ing
ca e ul conside a ion o in e ac ions wi h o he c ops o p e en ad e se impac s on soil
heal h and pes dynamics. The u iliza ion o di e si ied c op o a ions is seen as pi o al in
add essing sus ainabili y challenges in mode n ag icul u e. Howe e , designing o a ions
ha s ike comp omises ac oss mul iple sus ainabili y domains emains a pe sis en chal-
lenge [74]. Pes s and diseases pose ano he se o challenges, as esis an a ie ies o NUSs
may be limi ed, ende ing hem suscep ible o a ious h ea s. In his con ex , moni o ing
and managing pes s and diseases become challenging due o he ela i e no el y o NUS
Figu e 1. NUS c ops wi h he po en ial o cul i a ion in Medi e anean en i onmen s. Rep esen a i e
images o di e en nu ien - ich and s ess- esilien eme gen c ops o sus ainable and heal hy ood
in he Medi e anean egion, g own unde ield condi ions ac oss di e en egions o Spain. The
images depic he c ops in he ield (ou e images) and he panicle, in lo escence, o u le o each
c op (inne image). The c ops included in he igu e a e Chenopodium quinoa Willd., Ama an hus sp.,
Fagopy um sp., E ag os is e (Zucc.) T o e , Sal ia hispanica L., Mo inga olei e a Lam., Cannabis sa i a L.,
and mille (Pennise um glaucum (L.) R.B . Te ezia cla e yi Cha in (wi h Helian hemum alme iense Pau)).
Tole ance o hea has been epo ed o ama an h, a C4 plan , and mille [
42
,
57
],
while mille , ama an h, hemp, and quinoa can wi hs and ela i ely saline soils [
45
,
61
–
63
].
Buckwhea h i es in cold clima es and is well sui ed o poo soils, wi h educed wa e
equi emen s [
64
]. By con as , ce ain dese u le species om Te ezia sp. and Ti mania
sp. ace popula ion decline due o educed ain all and changes in land use, p omp ing
esea ch on domes ica ion o a d ough - esis an c op in line wi h sus ainable p ac ices [
33
].
O e all, hese mino c ops can h i e in ma ginal soils.
The p ima y challenge hinde ing hei expansion and in oduc ion in egions like
he Medi e anean basin is he sca ci y o comme cially a ailable a ie ies well adap ed
o he speci ic en i onmen al condi ions o hese a eas. Fo example, pho ope iod is he
p ima y obs acle o quinoa’s adap abili y in he no he n hemisphe e [
65
]. Quinoa is a
acul a i e sho -day and day-neu al plan [
66
]. Pho ope iod is also a majo limi a ion o
chia cul i a ion in Eu ope because i equi es a lo al induc ion pho ope iod o a ound 12 h
Plan s 2024,13, 1914 7 o 45
o da kness [
67
]. Fu he mo e, quinoa and buckwhea g owing unde high empe a u es
du ing lowe ing can expe ience d ama ic yield losses [
50
,
68
]. On he con a y, i has been
shown ha one o he p ima y cons ain s on he g ow h and p oduc i i y o mo inga is low
empe a u es. Mo inga h i es in opical and sub opical clima es. Mo inga can wi hs and
ligh os s down o
−
3
◦
C. Addi ionally, i is a demanding c op in e ms o ligh [
60
].
In he case o e , lodging ep esen s he mos signi ican issue limi ing yields [
69
], and
d ough adap a ion in dese u le myco hizal plan s is key when aiming a expanding
i s cul i a ion and in ol es enhancing plan nu i ion, pho osyn hesis, wa e use e iciency,
and oo coloniza ion mo phology [
41
]. Addi ionally, “ u micul u e,” an ag icul u al
app oach wi h minimal wa e inpu and sus ainable p ac ices, enables he cul i a ion o
his na u al esou ce in a id and semi-a id a eas [34].
The e o e, in he con ex o clima e change, ai s ha make eme ging c ops well sui ed
o Medi e anean condi ions include ole ance o he ha sh en i onmen al condi ions
expec ed o in ensi y in he coming yea s such as d ough , hea , and salini y [70].
Table 1. A lis o NUS c ops wi h he po en ial o cul i a ion in Medi e anean en i onmen s.
Common
Name Scien i ic Name Bo anical Family Type Abio ic S ess Tole ance
Cold Hea D ough Salini y
Quinoa Chenopodium quinoa Willd. Ama an haceae Dico yledons/He baceous/C3 Medium
Medium/low
High
Medium/high
Ama an h Ama an hus sp. (75 species
[71]) Ama an haceae Dico yledons/He baceous/C4
Medium/low
High High Medium
Buckwhea Fagopy um sp. Polygonaceae Dico yledons/He baceous/C3 Medium
Medium/low
High Medium
Mille /So ghum
Sub amilies:
- Chlo idoideae
- Panicoideae
(11 species excluding e )
[72]
Poaceae
Monoco yledons/He baceous/C4
Low High High High
Te E ag os is e (Zucc.)
T o e Poaceae
Monoco yledons/He baceous/C4
Medium High Medium
Medium/low
Chia Sal ia hispanica L. Lamiaceae Dico yledons/He baceous/C3 Low Medium Medium Medium
Mo inga Mo inga olei e a Lam. Mo ingaceae Dico yledons/Woody/C3 Ve y low High High
Medium/high
Hemp Cannabis sa i a L. Cannabaceae Dico yledons/He baceous/C3 Low
Medium/high
High High
Dese u les Te ezia cla e yi Cha in
(wi h Helian hemum
alme iense Pau)
Pezizaceae
(Cis aceae) Ascomyce e
(Dico yledons/woody/C3)
Medium/high
Medium Medium
Medium/low
2.4. Challenges and Limi a ions
The cul i a ion o NUSs poses a ious challenges ac oss di e en ace s o he ag icul-
u al and supply chain landscape. Ongoing h ea s o ag icul u al en i onmen s s em om
a g owing wo ld popula ion, limi ed a able land, and swi clima e changes. P ojec ions
sugges ha he global popula ion will su pass 9 billion by 2050 [
73
]. The challenge o
eeding his expanding popula ion is a global conce n. To secu e ood and ecosys em
s abili y, i is essen ial o de elop NUSs as c ops o sus ainable ag icul u e, emphasizing
inc eased ne p oduc ion while minimizing ad e se en i onmen al impac s. Ne e heless,
inco po a ing NUSs in o exis ing o a ions in oduces complexi ies, necessi a ing ca e ul
conside a ion o in e ac ions wi h o he c ops o p e en ad e se impac s on soil heal h and
pes dynamics. The u iliza ion o di e si ied c op o a ions is seen as pi o al in add essing
sus ainabili y challenges in mode n ag icul u e. Howe e , designing o a ions ha s ike
comp omises ac oss mul iple sus ainabili y domains emains a pe sis en challenge [
74
].
Pes s and diseases pose ano he se o challenges, as esis an a ie ies o NUSs may be
limi ed, ende ing hem suscep ible o a ious h ea s. In his con ex , moni o ing and
managing pes s and diseases become challenging due o he ela i e no el y o NUS in
ce ain egions, whe e exis ing con ol measu es may no be well es ablished. Ad ance-
men s in documen ing pa hogens and pes s a ec ing NUSs, along wi h he de elopmen o
co esponding managemen s a egies, a e, he e o e, essen ial [75].
NUSs ace an addi ional challenge due o he g owing in e es o consume s wo ldwide
in aligning hei ood consump ion wi h sus ainabili y [
76
] and he inc easing demand
o heal hie ood choices [
77
]. These a e he ac o s d i ing he inc easing in e es in
nu ien - ich and s ess- esilien NUS c ops. Fu he mo e, p ocessing eme ging c ops aces
Plan s 2024,13, 1914 8 o 45
in as uc u e and echnological challenges. I is c ucial o adap exis ing acili ies o he p o-
cessing equi emen s o NUSs and concu en ly es ablish quali y con ol measu es o mee
ma ke s anda ds. Despi e he g ea in e es in NUSs, mos a e no ye well adap ed and
highly ma ke able. De eloping e icien supply chains, encompassing logis ics, dis ibu-
ion, anspo a ion, s o age, and dis ibu ion ne wo ks, is c ucial [
78
]. Limi ed awa eness
among consume s and e aile s necessi a es ma ke ing and educa ional ini ia i es. The
speci ic niche ma ke s associa ed wi h NUSs c ea e challenges in es ablishing demand,
equi ing a ge ed ma ke ing s a egies. Socio-economic ac o s u he complica e he
adop ion o NUSs. Con incing a me s o emb ace change and cul i a e NUS c ops, espe-
cially when adi ional c ops ha e es ablished ma ke s and p o en ag onomic p ac ices, is
a hu dle. The economic easibili y o no el c ops
´
cul i a ion compa ed o adi ional c ops
plays a pi o al ole, equi ing assu ances o long- e m inancial iabili y. Addi ionally,
he lack o suppo i e policies, such as esea ch incen i es o inancial aid, can impede
widesp ead adop ion.
Thus, add essing all he abo e-men ioned challenges is essen ial o he success ul
in eg a ion o NUS c ops in o ag icul u al p ac ices, ensu ing sus ainable cul i a ion,
economic iabili y, and widesp ead adop ion. Collabo a ion among a me s, esea che s,
policymake s, and indus y s akeholde s is c ucial o de eloping sus ainable cul i a ion
p ac ices, imp o ing p ocessing echnologies, and c ea ing e ec i e ma ke s a egies.
Addi ionally, in es men s in esea ch and de elopmen , ex ension se ices, and capaci y-
building p og ams can con ibu e o he success ul in eg a ion o eme ging c ops in o
ag icul u al sys ems and ma ke s.
2.5. Gene ic and B eeding Ad ances
Gene ic imp o emen in NUS c ops has been conside ably lowe compa ed o majo
c ops. Consequen ly, he e exis s signi ican po en ial o enhancemen in a ious ag onom-
ical and nu i ional ai s. Addi ionally, many o hese c ops o igina e om egions wi h
ag oclima ic condi ions as ly di e en om hose in he Medi e anean. Hence, ongoing
b eeding e o s a e essen ial o de elop a ie ies be e adap ed o Medi e anean ag ocli-
ma ic condi ions. As p e iously men ioned, pho ope iod sensi i i y poses a challenge o
cul i a ing some NUS c ops in he Medi e anean basin. Nume ous accessions o quinoa,
ama an h, chia, buckwhea , and hemp equi e sho days o op imal p oduc ion. Fo
ins ance, quinoa exhibi s wo ge mplasm pools: he p ima y Andean highland quinoa
and he seconda y cen al and sou he n Chilean quinoa [
79
]. Fo una ely, ce ain quinoa
accessions showing day-leng h insensi i i y ha e been iden i ied in he coas al condi ions
o sou he n Chile. These ha e been u ilized o de elop a ie ies capable o h i ing in
Eu opean long-day en i onmen s. Eu opean a ie ies, de eloped in Denma k and he
Ne he lands, a e cu en ly a ailable, and a b eeding p og am is unde way in Spain o c ea e
new a ie ies speci ically adap ed o Medi e anean ield condi ions [
80
]. Simila ly, while
many buckwhea a ie ies a e sensi i e o pho ope iod, Eu ope now cul i a es a ie ies
ha a e no in luenced by day leng h [
81
]. Rega ding chia, he e a e a ew pa en ed a i-
e ies, such as he ‘Ou o’ a ie y om F ance, exhibi ing long-day lowe ing cha ac e is ics,
ob ained h ough selec i e b eeding [
82
]. Hemp, being a sho -day plan wi h lowe ing
highly dependen on pho ope iod and empe a u e, also includes a ie ies like ‘Finola’ ha
can ini ia e lowe ing independen ly o pho ope iod [83].
One o he key cons ain s o c op cul i a ion in he Medi e anean basin is he
occu ence o episodes o high empe a u es du ing he c op g owing cycle. Nume ous
s udies ha e demons a ed ha ele a ed empe a u es a lowe ing ime can signi ican ly
diminish yield in quinoa and buckwhea [
50
,
68
]. Consequen ly, enhancing hea ole ance
has become a c ucial b eeding a ge o inc ease he yield o quinoa and buckwhea in he
Medi e anean basin.
The success o de eloping new c op a ie ies ha h i e in di e en en i onmen s
and exhibi enhanced ag onomic and nu i ional ai s depends on gene ic a iabili y, un-
de s anding he gene ics o a ge ai s, and ha ing e ec i e imp o emen me hods and
Plan s 2024,13, 1914 9 o 45
ools. Quinoa ge mplasm exhibi s high di e si y, allowing i s adap a ion o a ious en i-
onmen s. Howe e , accessing Sou h Ame ican ge mplasm p esen s limi a ions, equi ing
nego ia ions wi h na ional go e nmen s and o en necessi a ing acili a ion by in e na ional
o ganiza ions like FAO [
84
]. Despi e being domes ica ed 7000 yea s ago by ancien Andean
ci iliza ions, quinoa b eeding p og ams only commenced in he 1960s in he Andes and
la e in o he egions om he 1970s onwa ds [
85
]. Se e al quinoa b eeding p og ams,
including hyb idiza ion, a e ongoing in di e en coun ies. Howe e , scien i ic knowledge
abou c i ical aspec s o quinoa b eeding, such as he inhe i ance o desi able ag onomic
and quali y ai s o he iden i ica ion o molecula ma ke s linked o hem o ma ke -
assis ed selec ion, emains limi ed. The sequencing o he quinoa genome by Ja is e al.
(2017) [
86
] has p o ided a powe ul ool o expedi e quinoa b eeding. Fo ins ance, his
genome sequence acili a ed he iden i ica ion o a gene con olling saponin con en [
86
].
Mo eo e , he a ailabili y o he quinoa genome sequence has enabled he iden i ica ion,
h ough genome-wide associa ion s udies (GWAS), o candida e genes o pho ope iodic
lowe ing egula ion, housand seed weigh , and o he ag onomic ai s [
87
,
88
]. Addi ion-
ally, ex eme g adien boos ing has been employed o iden i y candida e genes po en ially
con olling be alain con en in quinoa seeds [
89
]. These s udies open he possibili y o
u ilizing ma ke -assis ed selec ion in quinoa b eeding.
In con as , a no able sca ci y o gene ic di e si y has been obse ed wi hin domes-
ica ed chia accessions, wi h sligh ly g ea e a iabili y p esen in wild accessions [
82
].
Fu he mo e, gene banks hold limi ed chia ge mplasm [
90
]. Addi ional cons ain s o
chia b eeding include he p esence o small and agile co ollas, used lowe pa s, and a
p opensi y o sel -pollina ion, ac o s ha ha e impeded he es ablishmen o b eeding
p og ams cen ed on hyb ids [
91
]. Chia has no been he ocus o ex ensi e mode n plan
b eeding e o s, and imp o ed cul i a s ha e p ima ily a isen h ough he selec ion o
lines om mixed ge mplasm sou ces, ypically land aces [
91
]. Thus, limi ed knowledge
exis s ega ding he gene ics o pe inen ai s in chia, al hough some p og ess has been
achie ed. S udies on he inhe i ance o pho ope iod esponse, seed colou , lodging esis-
ance, and sha e ing esis ance ha e been conduc ed [
91
]. Also, ansc ip omic s udies
ha e iden i ied genes in ol ed in seconda y me aboli es and oil biosyn hesis [
92
]. Fu he -
mo e, chia genome sequencing has acili a ed he cha ac e iza ion o genes associa ed wi h
polyunsa u a ed a y acids (PUFAs), mucilage biosyn hesis, and hose coding o oleosin,
caleosin, and s e oleosin [93].
Ama an h is conside ed an o phan c op due o he limi ed e o s di ec ed owa ds
i s gene ic imp o emen [
94
]. Ne e heless, some p og ess has been achie ed h ough
hyb idiza ion (including hyb ids be ween di e en species), selec ion, and mu agenesis
me hods [
79
]. While mos a ie ies ha e been de eloped using con en ional b eeding,
gamma i adia ion-induced mu a ion has also been employed in ama an h o he de el-
opmen o new cul i a s [
95
]. None heless, gene ic s udies in ama an h a e sca ce, wi h
ma ke s iden i ied o plan g ow h, mo phology, seed cha ac e is ics, and lowe ing be-
ha iou . Inhe i ance s udies ha e been conduc ed o nu i ional ac o s, including s a ch
cha ac e is ics, he pe ispe m laye o g ain, seed p o ein con en , and male s e ili y [
79
]. A
Quan i a i e T ai Loci (QTL) o lowe colou was iden i ied in A. hypocond iacus L. [
96
].
Addi ionally, a ecen GWAS in Ama an hus icolo L. iden i ied ma ke - ai associa ions
associa ed wi h b anching index, in lo escence colou , pe iole pigmen a ion, and e minal
in lo escence shape and a i ude [
97
]. No ewo hy, genomic esou ces o ama an hs in-
clude he genomes o se e al Ama an h spp. ( he genomes o A. hypocond iacus L., A. icolo
L., A. palme i S. Wa s., and A. c uen us L. a e a ailable in he NCBI da ase ), genome-
wide SNPs and SSRs o A. cauda us L. and A. hypochond iacus L. and ‘popAma an h,’ a
popula ion gene ic genome b owse o g ain Ama an hus and hei wild ela i es [95].
Among all buckwhea species, only wo a e cul i a ed o g ain ood p oduc ion: com-
mon buckwhea (Fagopy um esculen um Moench) and a a y buckwhea (F. a a icum (L.)
Gae n). Common buckwhea has a wide dis ibu ion in Asia, Eu ope, Ame ica, and Aus-
ia, while a a y buckwhea is p ima ily g own in Asia [
98
]. Va ious b eeding p og ams
Plan s 2024,13, 1914 16 o 45
seeds showed a lowe ca bohyd a e con en (26–42%) (Table 2) compa ed o he o he
pseudoce eals. I should be no ed ha a lowe a e age ca bohyd a e con en in chia seeds
could be due o he high alues ob ained o all o he nu i ional pa ame e s. Thus, since
ca bohyd a e con en s o plan ood a e calcula ed by di e ence, an inc ease in p o ein, a ,
and mois u e con en s will ul ima ely a ec he ca bohyd a e con en s and con ibu e o
obse ed inc eases. So ghum and e g ains and mo inga lea es a e also ich sou ces o
ca bohyd a es, wi h con en s ound in he li e a u e anging om 63.7 o 80%, 70.8 o 73.8%,
and 29.8 o 51.6%, espec i ely (Table 2). Impo an ly, he main ca bohyd a e componen o
all c ops s udied is s a ch, cons i u ing a ound 50–70% o i [157,223].
3.2. Bioac i e Compounds
The widesp ead u iliza ion o pseudoce eals is a ibu ed o hei a ou able nu-
i ional alue and he p esence o bioac i e compounds. These compounds, no ably
la onoids, phenolic compounds, and pep ides, exhibi nume ous bene icial e ec s on
human heal h [
224
]. Howe e , he ex ac ion condi ions signi ican ly impac he quan i y o
bioac i e compounds ex ac ed (sol en , ex ac ion ime, and empe a u e), leading o sub-
s an ial a ia ions in esul s epo ed in he li e a u e. Consequen ly, his li e a u e e iew
e ains om p o iding speci ic alues o each o he compounds ound in di e en c ops.
3.2.1. Phenolics
Phenolic compounds, cha ac e ized by hyd oxyl g oups on a oma ic hyd oca bon
ings, cons i u e seconda y plan me aboli es. This compound amily includes phenolic
acids, la onoids, phenols, quinines, couma ins, lignans, phenylp opanoids, and xan-
hones [
225
], exis ing in bo h ee and bound o ms. These compounds, pa icula ly
la onoids, phenolic compounds, and pep ides, con e a ious heal h bene i s h ough
an ioxidan capaci y, sca enging ee adicals, and p e en ing oxida i e s ess [150,172].
Pseudoce eals, like quinoa, ama an h, and buckwhea , exhibi ich concen a ions o
bioac i e phy ochemicals. Quinoa, in pa icula , boas s la onoids such as que ce in and
kaemp e ol, con ibu ing o an ioxidan p ope ies and heal h bene i s [
200
,
226
]. Buck-
whea s ands ou as he iches sou ce o phenolic acids, p edominan ly ound in ee
o ms, in luencing an ioxidan ac i i y [227]. Chia seeds con ain signi ican phenolic com-
pounds, including a ious la onoids, bu also di e penoid phenolics such as osma inic
acid, bo h con ibu ing o he an ioxidan p ope ies o hese seeds [
224
,
228
]. So ghum,
uniquely ich in bioac i e cons i uen s, p esen s high concen a ions o phenolic com-
pounds such as e ulic and p o oca echuic acids, as well as la onoids like que ce in and
kaemp e ol
[229,230]
. Te g ains p ominen ly ea u e e ulic acid and la one de i a i es,
wi h addi ional phenolic compounds such as p o oca echuic and cinnamic acids [
231
].
Simple phenolic compounds and la onoids ha e been ex ensi ely documen ed in dese
u les, as well as in edible mush ooms. Howe e , he e a e con lic ing epo s ega d-
ing he abili y o hese ungi o ei he abso b o syn hesize hese compounds, especially
la onoids [195,232].
In summa y, high- alue seed lou s, including hose om buckwhea , quinoa, so ghum,
and e , exhibi ele a ed le els o phenolic subs ances and la onoids, su passing whole-
g ain ce eal lou s in an ioxidan and an i-in lamma o y e ec s [
233
,
234
]. Mo inga lea es
a e ich in phy ochemicals, wi h phenolic acids, la onoids, and an hocyanins con ibu ing
o hei nu i ional p o ile [235,236].
3.2.2. Phy os e ols
Phy os e ols, lipophilic compounds s uc u ally akin o choles e ol, play a i al ole
in choles e ol me abolism and exhibi po en ial heal h bene i s agains cance , diabe es,
hepa op o ec i e e ec s, and ca dio ascula diseases h ough hei choles e ol-lowe ing
cha ac e is ics [
150
]. In quinoa seeds, majo phy os e ols include
β
-si os e ol, campes-
e ol, and s igmas e ol, wi h highe quan i ies han hose ound in ba ley, ye, mille , and
co n [
180
,
237
]. Ama an h seeds also con ain hese phy os e ols, wi h
β
-si os e ol as he

Plan s 2024,13, 1914 17 o 45
p edominan compound [
238
]. In e es ingly, acco ding o [
239
],
β
-si os e ol, a signi ican
phy os e ol, may possess an i-diabe ic p ope ies.
Buckwhea g ains ea u e
β
-si os e ol as he p ima y phy os e ol, along wi h o h-
e s like campes e ol, a enas e ol, and D-7-s igmas e ol. Addi ionally, he in equen ly
obse ed cycloa enol was iden i ied in buckwhea samples [
240
]. Chia seeds simila ly
con ain phy os e ols such as campes e ol, s igmas anol, s igmas e ol, D5-a enas e ol, and
β
-si os e ol [
241
]. So ghum g ains encompass common s e ols, including
β
-si os e ol,
campes e ol, and s igmas e ol, posi ioning so ghum as a ich sou ce o phy os e ols com-
pa ed o ui s, ege ables, and o he ce eals [230].
While he e is limi ed e idence in he li e a u e o he p esence o phy os e ols in e
g ain, one s udy epo ed he exis ence o β-si os e ol and β-si os e ol-3-O-β-D-glucoside
in e g ains [
242
]. Also, mo inga lea es ha e eme ged as a ich sou ce o phy os e ols,
wi h
β
-si os e ol being he majo compound, ollowed by s igmas e ol, campes e ol, and
∆5-a enas e ol [154].
Fungal s e ols in dese u les emain inadequa ely in es iga ed. Some epo s sug-
ges ha he Te ezia genus p edominan ly con ains b assicas e ol (98%), wi h e gos e ol
cons i u ing less han 2%. Howe e , e gos e ol is a majo cons i uen in ungal mem-
b anes [243].
3.2.3. Be alains
Be alains, ni ogen-compound plan pigmen s, ha e ga ne ed a en ion in unc ional
oods due o hei bioac i e p ope ies, including an ioxidan and an i-in lamma o y e -
ec s [
244
,
245
]. Be alains demons a e supe io an ioxidan ac i i y compa ed o polyphe-
nols and a e exclusi e o plan s o he o de Ca yophyllales, including ce ain cul i a s o
he amily Ama an haceae [246].
Quinoa seeds de i e hei yellow, black, and ed colou s om be alains, making
quinoa a po en ial c op o be alain ex ac ion, al hough some a ie ies may lack measu -
able amoun s o be alains [
244
]. Be alains a e u he ca ego ized in o ed- iole be acyanins
and yellow-o ange be axan hins [
247
]. The inco po a ion o hese pigmen s in ood p od-
uc s is o icially ce i ied by he Eu opean Union (addi i e E-162) and he US FDA [
248
].
Ama an hine and isoama an hine a e wo be acyanins ound in ama an h, while ed and
black quinoa seeds con ain be acyanins such as be anin and isobe anin [
249
]. Some s udies
epo low amoun s o be alains in speci ic ama an h a ie ies [250].
3.3. An inu i ional Fac o s
Ce ain phy ochemical compounds ound in seeds, such as phy a es, saponins, lec ins,
and p o ease inhibi o s, can exe bo h posi i e and nega i e e ec s on human heal h. A
low concen a ions, hese compounds may con ibu e o well-being by exhibi ing an ioxi-
dan p ope ies, an i-in lamma o y e ec s, and an i-cance o immune-enhancing e ec s.
Howe e , when p esen in highe concen a ions, hese phy ochemicals can in e e e wi h
he abso p ion o essen ial mine als, induce haemolysis, dis up p o ein diges ion and
abso p ion, o ha m he gu wall, po en ially leading o inc eased in es inal pe meabili y.
While a mode a e in ake o hese compounds o e s heal h bene i s, excessi e consump ion
should be a oided o main ain a balanced die and ensu e op imal nu ien abso p ion.
Addi ionally, oxala es and ni a es, na u ally occu ing compounds in a ious plan oods,
a e gene ally conside ed ha mless when consumed in mode a ion. O e all, a ious an inu-
i ional ac o s a e p esen in he edible pa s o NUS c ops, and hei in luence should
be conside ed du ing he assessmen o nu i ional cha ac e is ics. In his ega d, i is
c ucial o highligh he e y limi ed knowledge abou he p esence o an inu ien s in
he majo i y o he NUS c ops analysed in his wo k. This unde sco es he necessi y o
conduc ing addi ional esea ch on his opic. None heless, a concise analysis is p esen ed
he ein, and he indings a e summa ized in Table 3. Fu he mo e, o mi iga e he p esence
o an inu ien compounds, a ious ac ions o s a egies can be employed du ing ood
p ocessing, as ou lined in Table 3.
Plan s 2024,13, 1914 18 o 45
In he case o phy ic acid, also known as myo-inosi ol hexakisphospha e, i is com-
monly ound in g ains and seeds as phy a es. While phy ic acid in ake has epo ed bene i s
such as an ioxidan unc ion and he p e en ion o hea diseases and cance , i is no o ious
o inhibi ing he abso p ion o mine als, p o eins, and ace elemen s [
251
,
252
]. The o -
ma ion o insoluble complexes in he gas oin es inal ac ende s hem non-abso bable,
while enzyma ic deg ada ion o phy ic acid by phy ases imp o es he nu i ional alue o
ce eal, pseudoce eal, legume, and oilseed p oduc s [253–255].
Saponins, cha ac e ized by a linea a angemen o hexose o pen ose glycoside uni s
linked o sapogenin aglycones, o m complexes wi h p o eins, lipids, zinc, and i on, exhibi -
ing a haemoly ic e ec , albei abso bed in limi ed quan i ies; hey can enhance memb ane
pe meabili y, po en ially aiding in in es inal ood in ake and d ug assimila ion [
256
,
257
].
Howe e , hei bi e as e poses a challenge o he widesp ead u iliza ion o quinoa o
ama an h seeds [258].
Ano he an inu ien compound amily, p o ease inhibi o s, abundan in legumes,
ce eals, pseudoce eals, and oilseeds, o ms s able complexes wi h p o eoly ic enzymes,
in luencing human physiology in bo h bene icial and de imen al ways [
259
,
260
]. While
hey a e conside ed an inu i ional ac o s due o hei inhibi o y e ec s on diges i e en-
zymes, quinoa and ama an h seeds con ain minimal amoun s, unlike commonly consumed
g ains [
256
]. Con e sely, he limi ed diges ibili y o buckwhea p o ein is a ibu ed o
a ious ac o s, including p o ease inhibi o s, annins, and
α
-amylase inhibi o s [
162
]. Ox-
ala es, ound in a ious plan issues including lea es, s ems, hypoco yl– oo , and seeds,
hinde he abso p ion o calcium and magnesium, con ibu ing o kidney s one o ma ion.
Ama an h and quinoa a e conside ed high oxala e sou ces, wi h he highes concen a ions
obse ed in lea es and s ems. Howe e , he majo i y o oxala es a e ound in an insoluble
o m, and due o hei high concen a ions o calcium and magnesium, his may esul
in low oxala e abso babili y [
261
]. None heless, o mi iga e he isk o s one o ma ion,
limi ing oxala e consump ion o below 100 mg pe day is ad isable, wi h indi iduals p one
o kidney s ones po en ially equi ing u he educ ion o less han 50 mg pe day [
262
].
Cooking g ains be o e consump ion is essen ial o educe soluble oxala e in ake, as soluble
sou ces a e mo e eadily abso bed, posing a isk o kidney s one de elopmen [
263
]. In he
case o lec ins, whose oxici y is con ingen upon he quan i y consumed and he equency
o inges ion, hey a e p o eins ha can bind o cells, leading o s uc u al and physiological
damage in issues upon consump ion, esul ing in po en ially ad e se heal h e ec s [
264
].
They ha e been ound in quinoa, ama an h, o dese u le [265–267].
Ni a es, while no inhe en ly ha m ul, can con e o ni i es du ing diges ion, po-
en ially leading o he o ma ion o ni osamines in he s omach [
268
]. The Wo ld Heal h
O ganiza ion (WHO) has no speci ied a ecommended daily allowance (RDA) o ni a es,
bu he Eu opean Food Sa e y Au ho i y (EFSA) has se an accep able daily in ake (ADI) a
3.7 mg/kg o body weigh /day, app oxima ely 260 mg/day o a 70 kg adul . Gene ally,
g ains o seeds om selec ed NUS c ops all below his h eshold. Howe e , gi en he lack
o in o ma ion on ni a e con en in he g een pa s o hese c ops, i is ad isable o conduc
ni a e con en analyses when consumed as ege ables, as ege ables a e a p ima y sou ce
o ni a es in plan -based oods.
Plan s 2024,13, 1914 19 o 45
Table 3. Mainly an inu i ional ac o s in selec ed c ops and he ea men s o inhibi hei ac ion.
An inu ien
Fac o Phy ic Acid Saponins P o ease
Inhibi o s Lec ins Oxala es Ni a es Re e ences
C op and Usual
T ea men g/100 g mg/100 g mg/100 g g/100 g mg/100 g mg/100 g
Quinoa 0.50–1.39 1.90–6.24; 199.07
629.7–1633.3 ** 2.4 * g/kg 5.2 HU’; 9.3% 4.31; 6.17–9.45 21.83 [158,258,266,269]
Ama an h 0.61–1.34 5.33–10.66 HU/mg
79–186 0.519–1.016
TIU/mg
0.5–7.3;
11.2–130.6
HAU/mg
178–278;
46–152 48–84 [258,261,267,270,
271]
Buckwhea 0.63; 1.7 0.058 55 αU/mg p o ein;
94 αU/mg p o ein NYR (+) 93–155 NYR (−) [161,272–274]
Mille 0.12–0.92; 0.336–0.649;
0.519–0.784; 0.64–0.81 0.039–0.044 0.385–0.985
TIU/mg lou NYR (−) 11.3–20.0 NYR (−) [164,272,275–277]
Te 0.682–1.374; 1.35 722 NYR NYR (−) 3.50–5.27 *** NYR (−) [278–281]
Chia 1.5–2.6 616 445 TIU/mg p o ein NYR NYR (−) NYR (−) [163,282]
Mo inga (seeds) 0.115 δ;2.54 1080 δ15.4; 120 δNYR (+) 2.21 δNYR [283,284]
Hemp 2.66–3.08 123.54 α3.6% βNYR NYR NYR [285–287]
Desse u les 0.581 12.38 NYR 18.11 HU NYR NYR [265,288]
Usual T ea men
o diminish he
an inu ien
ac o
concen a ion o
deac i a ion.
In gene al,
an inu ien
le els can be
con olled by
cul i a selec ion
and b eeding,
ield
managemen ,
and p ocessing
Soaking, S eeping,
Mal ing,
Fe men a ion,
Ge mina ion,
Ex usion, Popped
g ains,
Phy ase ac i a ion by
sp ou ing,
Ge mina ion, o
lowe ing he pH
Inclusion o
exogenous phy ases
Ag onomic condi ions
Gene ic enginee ing
Ge mina ion
Popped seeds
Ex usion
Swee a ie ies
(<110 mg/100 g)
Ag onomic
condi ions
Cooking a ound
100 ◦C o a
highe
empe a u es
Popped g ains
Ex usion
Cooking a ound
100 ◦C o mo e
Soaking
Ex usion
Popped seeds
Ge mina ion
Wa e
cooking-soluble
oxala e, Soak-
ing/S eeping
Con ol o e il-
iza ion/soil
Wa e cooking
Soaking/S eeping
Con ol o e il-
iza ion/soil
* g/kg: he amoun o enzyme inhibi ed pe kg seed meal; HU’: Haemagglu ina ing Uni s
×
10
−6
pe kg o
seed meal, ea ed a blood cells; TIU/mg, HAU/mg, and HU/mg: T ypsin Inhibi o y Uni s, Haemoly ic
Ac i i y Uni s, and Haemoly ic Uni s pe mg o he sample on a d y weigh basis. ** exp essed in sapogenins
( he hyd olysed p oduc o saponins); *** he amoun in s aw;
δ
de a ed seed lou ;
α
in he ex ac ;
β
ypsin
inhibi o ; NYR no ye epo ed; p esence: (+); appa en ly a a lowe amoun han o he g ains/seeds: (−).
3.4. Nu i ional Challenges and Conside a ions
Cu en ly, he global communi y con on s a spec um o nu i ional challenges en-
compassing malnu i ion and hunge , jux aposed wi h conce ns ega ding obesi y and
die - ela ed ailmen s such as diabe es and ca dio ascula diseases. Concu en ly, he exi-
gencies o clima e change and wa e sca ci y necessi a e a en ion, alongside unde u ilized
ood was es, byp oduc s, and he inadequacy in mee ing he p o ein demand o he wo ld
popula ion [
289
]. In his ega d, i becomes necessa y, and almos manda o y, o le e age
and capi alize on he scien i ic and cul u al esou ces and knowledge a ou disposal.
Hence, aligning wi h he cen al heme o his e iew, he possibili ies p esen ed by he
u iliza ion and e alua ion o NUSs a e well ecognized. These possibili ies con ibu e o
enhanced biodi e si y and sus ainable de elopmen , no only a he ag onomic le el bu
also wi hin he indus ial con ex . The ul ima e goal is o achie e imp o ed p oduc ion,
enhanced nu i ion, a heal hie en i onmen , and an o e all be e quali y o li e o all,
lea ing no one behind [290].
Beyond popula labels such as ‘supe oods’ o ‘mi acle seeds o ees’, he c ops
discussed in his wo k exhibi a chemical composi ion in mac o- and mic onu ien s ha
a e ei he simila o o supe io o he ou main plan c ops domina ing he global ag i ood
sys em—namely, ice, whea , soybeans, and co n. No ably, he p oduc ion o hese majo
c ops is concen a ed in a ew coun ies globally (USA, China, India, A gen ina, B azil, and
Russia). A dis inc i e ad an age o many eme ging c ops highligh ed in his s udy is hei
supe io adap abili y o abio ic s ess condi ions, as de ailed in Sec ion 2on ag onomical
Plan s 2024,13, 1914 20 o 45
aspec s ela ed o NUSs o he Medi e anean egion [
10
,
291
]. Hence, i is impe a i e o
unde ake mo e comp ehensi e s udies on he in oduc ion and ad ancemen o hese
eme ging c ops, in ol ing gene ic imp o emen , hyb idiza ion, a i icial in elligence, and
enhanced con ol o e abio ic s esses and pes s. The objec i e is o di e si y bo h he
plan aw ma e ials dedica ed o ood and he na ional and local p oduc ion le els. This
s a egic app oach aims o mi iga e ecen ood c ises, such as hose expe ienced du ing
he COVID-19 pandemic o con lic s in egions like Uk aine and Gaza. The acquisi ion
o g ea e ag onomic knowledge will empowe go e nmen s o implemen ag icul u al
p og ams and policies ha posi i ely impac nu i ion.
Conduc ing esea ch and dissemina ing indings on he nu i ional aspec s o eme ging
c ops, including ood composi ion, die a y assessmen s, and human nu i ional equi e-
men s, is essen ial o in eg a ing hese c ops in o socie y and highligh ing hei heal h
bene i s. This e o add esses he c ucial nu i ional challenge o mee ing he ising global
p o ein demand h ough sus ainable p oduc ion, di e si ied p o ein sou ces, and e icien
esou ce use.
As s a ed in Table 2, he seed p o ein con en shown by some can exceed 20% (e.g.,
chia [
150
] o hemp [
292
]). Fu he mo e, all exhibi highe p o ein con en s han commonly
consumed ce eals [
145
,
171
,
176
]. In his con ex , an unde u ilized and aluable esou ce o
be e alued and exploi ed is he p o ein con ibu ion om o he pa s o he plan besides
he seeds, no ably he lea es. C ops like mo inga, o ins ance, p esen p o ein alues
anging be ween 16 and 40% in his ega d [
166
] o dese u les, wi h alues anging
be ween 14 and 29% [
178
,
195
]. The p o ein con en can be u he enhanced h ough he
ex ac ion and comme cializa ion o o he componen s applicable in he ood indus y, such
as oil, s a ch, o ib e ac ion. This p ocess esul s in p o ein concen a es o isola es [
293
],
which can se e as plan -based ing edien s in p oduc s wi h high p o ein con en ca e ing
o ulne able popula ions such as he elde ly, malnou ished child en, o p egnan women.
Recen ad ances in ood and heal h emphasize he po en ial o pep ides, mono- and
polyunsa u a ed a y acids, pigmen s, and low glycaemic index s a ches om hese c ops,
sugges ing p omising esea ch di ec ions. This p og ess equi es enhancing and upda ing
ex ac ion echnologies, such as mic owa es, ul asound, high p essu es, e men a ion,
ibo-elec os a ics, and supe c i ical luids. Addi ionally, he e is ongoing explo a ion
and u iliza ion o ex ac ion me hodologies, such as hose assis ed by enzymes o eu ec ic
sol en s [294–299].
Ano he signi ican nu i ional challenge associa ed wi h hese c ops is he u iliza ion
o e alua ion o an inu i ional compounds, pa icula ly ound in he seeds. Examples, as
desc ibed ea lie in his e iew, include saponins in quinoa [
300
], annins in so ghum [
301
],
o phy ic acid in ama an h, chia, and mille [
302
]. Ini ially u ilized by he plan as a de-
ence o su i al mechanism agains pes s, p eda o s, o ad e se wea he condi ions, hese
componen s we e adi ionally deemed exclusi ely an inu i ional o decades. Howe e ,
ecen scien i ic s udies ha e shi ed he ocus, a ibu ing ce ain heal h bene i s a low
concen a ions, such as an ioxidan o an i-in lamma o y p ope ies [
303
–
305
]. The de-
elopmen o a mo e iendly and p o i able echnology wi h hese aw ma e ials a he
indus ial le el will inc ease his e alua ion and help o enhance he bene i s o hese c ops
a he ood le el.
3.5. Technological Aspec s
The su ge in quinoa demand is a ibu ed o i s dis inc i e nu i ional and echno-
unc ional cha ac e is ics. Fo me ly s igma ized as a symbol o u al po e y and indigene-
i y, quinoa has gained supe ood s a us [
306
]. No ably, quinoa s a ch, wi h i s high peak
iscosi y, holds p omise o enhancing echno- unc ionali y in glu en- ee dough o mula-
ions and ein o cing edible biodeg adable ilms. Resea ch by Co doba-Ce ón e al. [
307
]
showcased imp o ed echno- unc ional p ope ies in p o ein- ich pas a when inco po a ing
de a ed high-p o ein quinoa lou . Fungal e men a ion has eme ged as a iable me hod o
enhance he bioac i i y o lou s, as demons a ed in quinoa, leading o inc eased le els o
Plan s 2024,13, 1914 21 o 45
esis an s a ch and ib e [
308
]. Ama an h p o ein su passes ce eals in biological alue and
exhibi s a ou able echno- unc ional p ope ies, including high solubili y and oaming
capaci y. S udies by Kie ul e al. [
309
] demons a ed ha ama an h and quinoa lou s,
e aining a po ion o hei p o ein con en , possess excellen echno- unc ional p ope ies,
quali ying hem as na u al al e na i e su ac an s o s abilizing ood emulsions [309].
Chia seeds, endowed wi h in insic echnological cha ac e is ics such as high wa e -
holding capaci y and obus gelling p ope ies, a e sui able as a eplace s in baked and
p ocessed mea p oduc s [
310
,
311
]. While o he NUSs hold nu i ional in e es in ood
o mula ion, hei echnological applica ions p esen challenges. Hemp seed p o ein (o
app oxima ely 20–25%), cha ac e ized by low alle genici y and a comple e amino acid
p o ile, aces limi a ions in echno- unc ional p ope ies o success ul o mula ion in
ood p oduc s [
312
]. Ex usion echnology has p o en success ul in o e coming hese
limi a ions, pa icula ly o p oduc s mimicking mea [
313
]. Mo inga encoun e s obs acles
in ood o mula ion due o o ganolep ic cha ac e is ics such as an as ingen bi e as e
and unpleasan smell. Encapsula ion echnology has been p oposed o add ess hese
challenges while enhancing he mal s abili y [
314
]. Al e na i ely, he use o mo inga lowe
buds in powde ed o ma has been sugges ed o di ec inco po a ion in o pas a p oduc s,
enhancing nu i ional p o iles and an ioxidan ac i i y [167].
Dese u le (Te ezia a ena ia), al hough no lis ed as an edible ood in he Eu opean
Union, p esen s an in iguing oppo uni y o sus ainable plan -based mea analogues due
o i s unique a oma and composi ion analogous o mea [
315
]. Ne e heless, dese u les
ha e been success ully employed o enhance he nu i ional composi ion and an ioxidan
con en o biscui s [169].
3.5.1. Culina y and P ocessing Aspec s
The comp ehensi e examina ion o he culina y and p ocessing cha ac e is ics o
nu ien - ich and s ess- esilien eme gen c ops p o ides insigh s in o hei po en ial
applica ions in adi ional cuisines and hei nu i ional bene i s o p omo ing hei con-
sump ion in con en ional culina y p ac ices and imp o ing nu aceu icals o high-quali y
unc ional oods [
316
]. Quinoa, o example, is ex ensi ely u ilized as a g ain o lou in
he p epa a ion o b ead, soups, and cooked p oduc s, ei he independen ly o in con-
junc ion wi h o he lou s [
54
]. Gi en i s compa ibili y wi h adi ional cuisine cul u e,
quinoa seamlessly in eg a es in o Sou h Ame ican dishes, lending i s nu y la ou and
lu y ex u e o adi ional Pe u ian quinoa s ews and con empo a y salads, showcasing
adap abili y ac oss di e se global cuisines [
317
]. Mo eo e , expanded o pu ed quinoa
in oduces ano he a ia ion o adi ional quinoa p oduc s. This in ol es subjec ing he
g ain o a high empe a u e and p essu e, esul ing in a quinoa pop. High empe a u es and
long p ocessing imes, common in ene gy ba s and ins an ce eal p oduc ion, can cause a
dec ease in p o ein and linoleic acids, impac ing he nu i ional quali y o he p oduc [
318
].
Chia seeds s and ou o hei la ou and imp essi e wa e -abso bing p ope ies,
o e ing a e sa ile, gel-like consis ency ha makes hem a a ou ed ing edien in be e ages,
puddings, smoo hies, and a ious swee and sa ou y ecipes o a nu ien - ich boos [
319
].
Thei combined nu i ional and echnological a ibu es a e op imal o use in baked goods
as an egg eplacemen ing edien , o in he p oduc ion o glu en- ee b ead o pas a, owing
o hei hyd ocolloid and hickening p ope ies [
320
]. Nume ous s udies ha e explo ed
he inco po a ion o chia seeds and chia mucilage gel as subs i u es o eggs, oil, o a in
di e se ood p oduc s esul ing in mo e nu i ious i ems wi h accep able senso y quali ies,
especially e iden in pound cakes, educed- a b ead baking, and imp o ed quali y in swee
whea doughs [
321
–
323
]. Addi ionally, plan -based mea analogues o e an inno a i e
s a egy o emula ing he senso y and nu i ional a ibu es o adi ional mea p oduc s.
Wi h an inc easing demand o hese al e na i es, he e is a g owing explo a ion o aw
ma e ials capable o eplica ing mea p oduc s. Chia seeds eme ge as a p omising candida e.
De a ed chia lou , known o i s high p o ein con en , is aluable, especially due o he
chia mucilage ac ion. This can se e as an e ec i e ing edien con ibu ing o enhancing

Plan s 2024,13, 1914 22 o 45
he ex u al p ope ies o plan -based mea al e na i es [
324
]. Quinoa and chia, as iable
op ions o ein eg a ing in o he ood alue chain, o e cop oduc s ha , when u ilized in
mea p oduc ion inno a ion, especially in con olling lipid oxida ion, can play a c ucial
ole in enhancing shel li e [325,326].
Ama an h, wi h i s dis inc i e ea hy and peppe y la ou p o ile, has become a
culina y co ne s one in Cen al and Sou h Ame ica, India, and A ica. In Mexico, i
is a ou ed o c a ing delec able, swee ea s, while in India, i ea u es a my iad o
bo h sa ou y and swee dishes. I s minuscule g ains con ibu e a unique ex u e and
nu i ional ichness o a di e se ange o culina y c ea ions, including soups, s ews, and
baked goods. Recen ly, he e has been a esu gence in he popula i y o ama an h, ma king
a e i al i e cen u ies a e i se ed as a s aple ood o ancien Mesoame icans [
327
].
Today, ama an h is easily accessible in a a ie y o comme cial ood p oduc s wo ldwide,
showcasing i s endu ing popula i y and ecognized nu i ional bene i s [
328
]. Recen
de elopmen s in ol e in eg a ing g ain ama an h in o ene gy oods, including b eak as
i ems, b ead, c acke s, and pancake mixes. Addi ionally, i can be enjoyed popped as a
wholesome snack op ion [327].
The scien i ic li e a u e obus ly suppo s he culina y u iliza ion o Mo inga olei e a,
highligh ing i s po en ial as a aluable sou ce o phy ochemicals wi h di e se applica-
ions in unc ional and medicinal oods [
329
–
332
]. The inco po a ion o mo inga in o
bake y p oduc s has shown signi ican nu i ional bene i s, such as imp o ed diges ibili y,
enhanced dough s abili y, inc eased an ioxidan capaci y, and ex ended p ese a ion [
333
].
Addi ionally, mo inga’s an ioxidan and an imic obial p ope ies make i sui able o
educing lipid oxida ion and inhibi ing bac e ial g ow h, he eby enhancing he nu i ional
con en o ood i ems like ish o chicken bu ge s, c eam cheese, o mu on pa ies [
334
–
336
].
P o ein ex ac s de i ed om ac ions o mo inga seeds ha e been epo ed as a po en ial
ing edien o uphold ex u e and educe syne esis in yoghu p oduc ion [
337
]. As he end
in ood o i ica ion is inc easing, mo inga has been explo ed as a unc ional ing edien
in o i ica ion in p o ein and ib e wi h accep able senso y quali y in p oduc s such as
pas a, cookies, b ead, o empeh-like e men ed p oduc s [
338
–
340
]. Inco po a ing mo inga
in o he ood indus y o e s a signi ican oppo uni y o ackle nu i ional de iciencies,
pa icula ly in egions wi h ex eme po e y o ood sca ci y [
341
]. By ha nessing mo inga’s
nu i ional ichness, ood manu ac u e s can de elop p oduc s ha p o ide nou ishmen
and enhance he well-being o communi ies in need [342].
Also, buckwhea , mille , and e a e no ewo hy eme ging c ops, pa icula ly as glu en-
ee g ains, making hem sui able o ood indus ial applica ions ha p io i ize nu i ious
and glu en- ee ood p oduc s [
343
]. Buckwhea is expe iencing inc eased popula i y in
Eas e n Eu ope, ea u ing in dishes like blini, and Asian cuisine h ough soba noodles.
I s nu i ional alue and p esence o bioac i e compounds like pep ides, la onoids, and
phenolics posi ion i as a p omising c op o de eloping glu en- ee p oduc s such as cook-
ies, b ead, o ocaccia o ca e o indi iduals wi h celiac disease [
344
,
345
]. In Cen al Asia,
whe e buckwhea is a die a y s aple, a s a egy o dec ease mea consump ion in ol es
in oducing 3D ood p in ing in he o mula ion o glu en- ee, plan -based mea al e -
na i es c ea ing and cus omizing balanced oods [
346
]. Aligned wi h buckwhea , mille ,
a die a y s aple in many A ican and Asian na ions, is u ilized in po idges, la b eads,
and pila s, and i s glu en- ee na u e also makes i a sui able al e na i e o indi iduals
wi h glu en sensi i i ies [
345
,
347
]. Also, he inc easing global demand o e men ed oods
has opened he possibili y o u ilizing mul ig ain mille , gi en i s subs an ial p obio ic
p ope ies [
348
]. Fo ins ance, e men ing whey ce eal be e ages wi h mille and mo h
bean p oduced an inno a i e dai y al e na i e wi h desi able unc ional and senso y ai s,
main aining accep abili y o 12 days [
349
]. O igina ing om E hiopia, e lou enhances
he echnological p ocess and quali y o whea b ead, boos ing i s nu i ional alue and
speci ic quali ies by imp o ing s uc u al and mechanical pa ame e s [
350
]. Addi ion-
ally, e se es as a subs i u e o b eadc umbs and soybean meals in con en ional mea
p oduc s, enhancing mois u e e en ion and senso y cha ac e is ics, making i sui able o
Plan s 2024,13, 1914 23 o 45
indi iduals wi h celiac disease and soy p o ein sensi i i y [
351
]. Fu he mo e, ex usion
cooking has been in es iga ed o c ea e expanded e -based p oduc s mixed wi h chickpea
lou , showing p omising physical, unc ional, and senso y a ibu es, indica ing po en ial
applica ions in he ood indus y [352].
Hemp seeds, known o hei nu y la ou , play a p ominen ole in global cuisine,
whe he sp inkled o e salads, blended in o smoo hies, o inco po a ed in o baked goods
con ibu ing o a boos in p o ein and omega-3 a y acids. Va ious p ocesses o p oducing
alue-added ing edien s om hemp seeds, including oil and p o ein, a e well sui ed o
a ge ed ood applica ions, p esen ing ma ke -led oppo uni ies. Examples include hemp
lou and de i ed oil o o i ied ood p oduc s, as well as he ex ac ion o p o ein isola es
and concen a es o plan -based mea and dai y al e na i es [
353
]. Fo ins ance, inco po-
a ing hemp lou in cookies, which impa s high an ioxidan ac i i y, o subs i u ing s a ch
wi h 20% hemp p o ein concen a e o enhance he quali y o glu en- ee b ead
[354,355]
o
e men ing hemp seeds o p oduce e men ed d inks wi h p ebio ic ac i i y, has esul ed in
an al e na i e dai y p oduc [
356
]. Fu he mo e, bo h d y and high-mois u e ex usion ech-
niques ha e esul ed in hemp-based p oduc s, including ene gy ba s and mea analogue
o mula ions, exhibi ing high senso y accep abili y [313,357].
Finally, in Middle Eas e n cuisine, dese u les a e e e ed as a delicacy, impa ing a
dis inc i e ea hy and umami la ou o dishes. They a e sa ou ed h ough a ious p epa-
a ions, including ying, boiling, o as a mea eplacemen in cooked dishes [
155
]. F om
ice-based delicacies o sauces and desse s, he adap abili y o dese u les showcases
hei endu ing appeal in bo h adi ional and con empo a y culina y se ings [
315
]. Due o
he nu i ional p o ile o his c op, which encompasses a wide ange o bioac i e chemical
cons i uen s, i has been u ilized as a he apeu ic sou ce [358].
3.5.2. Technological T ea men s
Despi e hei inc easing use in he ood indus y, mino ce eals and pseudoce eals o -
en ha e poo unc ional p ope ies o ce ain ood manu ac u ing p ocesses. To o e come
hese sho comings, lou s o g ains o hese species ha e been subjec ed o a ious echno-
logical ea men s ha induce s uc u al modi ica ions o hei main biopolyme s (s a ch
and p o eins), leading o signi ican imp o emen s in hei echno- unc ional p ope ies.
Among he echnological ea men s applied, he physical modi ica ion o s a ches, lou s,
and g ains has become inc easingly impo an , as i allows he unc ional imp o emen
o hese ma ices while main aining he s a us o ood ing edien s so ha hey can be
inco po a ed in o he o mula ion o clean label oods.
Hea Mois u e T ea men (HMT) is one o he mos commonly used and is ca ied ou
a empe a u es abo e he s a ch gela iniza ion empe a u e while main aining a mois u e
con en o less han 35%. HMT has been applied o buckwhea s a ch and lou o imp o e
hei he mal s abili y and nu i ional p ope ies, making hem mo e sui able o pas a and
baked p oduc s [
359
,
360
]. The ea men o quinoa wi h HMT has also been p oposed o
modi y he gela iniza ion beha iou , ela i e c ys allini y, and Fou ie T ans o m In a ed
(FTIR) spec a o s a ch, imp o ing i s ange o applica ions as a ood hickene [
361
], and
o inc ease s a ch/p o ein in e ac ion in lou s, he eby educing he es ima ed glycaemic
index [
362
]. Mic owa e-assis ed hyd o he mal ea men (MWT) has eme ged as a p omis-
ing al e na i e, as i allows o as e ea men and educes he amoun o ene gy equi ed
in compa ison o con en ional he mal hea ing. S udies on buckwhea g ains ha e shown
ha he mois u e con en du ing MWT modula es he inal p ope ies o he lou , hus
widening he ange o applica ions o buckwhea in he ood indus y, such as soups
and sauces, cakes, ozen desse s, and bake y p oduc s [
363
]. The use o MWT- ea ed
buckwhea lou in he p oduc ion o glu en- ee p oduc s esul ed in imp o ed dough
iscoelas ici y and b ead-speci ic olume, delayed s aling, and modula ed glucose elease
kine ics [
364
]. In he same way, MWT is e ec i e in modi ying he unc ional p ope ies
o quinoa lou s by al e ing hei p o ein and s a ch componen s, hus in luencing he
heological beha iou o b ead doughs, which a ec s b eadmaking pe o mance [365].
Plan s 2024,13, 1914 24 o 45
Annealing (ANN) is an al e na i e hyd o he mal ea men in which s a chy ma e ials
a e ea ed in excess wa e be ween he glass ansi ion and gela iniza ion empe a u es.
This echnology has been success ully used o imp o e he he mal s abili y o buckwhea
s a ch while modi ying i s
in i o
diges ibili y, inc easing slow diges ible s a ch and esis-
an s a ch [
366
]. P e ious s udies on he ANN ea men o ama an h s a ch ha e shown
ha he o ma ion o c oss-links be ween s a ch chains (amylose–amylopec in) al e s he
physicochemical p ope ies o ama an h s a ch wi h imp o ed he mal s abili y, inc easing
i s po en ial use as an ing edien in he ood indus y [367].
High-in ensi y ul asound (US) has also been used o he ea men o s a ches and
lou s, which a e gene ally suspended in an aqueous medium in which hey unde go
physical modi ica ion by he phenomenon o ca i a ion. This echnology has been applied
o mino ce eals such as cana y seeds o imp o e he unc ional p ope ies o he lou s [
368
].
Sonica ed de a ed cana y seed lou s p o ed o be e sa ile ing edien s wi h desi able
su ac an p ope ies o a wide ange o applica ions such as cakes, mu ins, and b ead
whe e ex u e enhancemen , s abili y, and consis ency a e c i ical.
High hyd os a ic p essu e (HHP) is a echnology gene ally used in he ea men and
p ese a ion o oods u s, which has ecen ly been applied o he ea men o ce eals and
pseudoce eals due o i s abili y o exe signi ican e ec s on s a ch and p o ein polyme s.
The abili y o modi y he hyd a ion and pas ing p ope ies o quinoa and buckwhea
s a ches, as well as he ex u e and heology o he gels o med om hem, has ecen ly
been in es iga ed [
369
–
371
]. HHP ea men s ha e also been applied o e and buckwhea
lou s [
372
] and whole buckwhea g ains [
373
], imp o ing he phenolic con en o he
esul ing lou s and hei s uc u ing capaci y in glu en- ee sys ems, depending on he
ea men a iables.
4. The Heal h Bene i s o NUSs o he Medi e anean Region
Globally, de eloped coun ies a e expe iencing an inc ease in he incidence o majo
non-communicable diseases such as ca dio ascula disease, diabe es, and obesi y. This
inc ease has led o a signi ican ise in mo bi-mo ali y and heal hca e expendi u e [
374
].
The sea ch o no el oods ha no only p o ide balanced nu i ion bu also ha e mul i-
ple bene icial heal h e ec s has made he s udy o unc ional oods an incipien ield. In
his con ex , he heal h-p omo ing aspec s o bioac i e compounds ound in NUS, such
as quinoa, chia, ama an h, hemp, mo inga, so ghum, buckwhea , e , o desse u les,
ha e been elucida ed. Quinoa and ama an h, in pa icula , no only ha e commendable
nu i ional p o iles bu a e also signi ican ly iche in bo h mic o- and phy onu ien s han
con en ional g ains, which has led o a g owing in e es in hese c ops o po en ial in eg a-
ion in o unc ional oods and nu aceu icals [
375
]. Recen s udies ha e ocused on quinoa
and ama an h in ending o iden i y nume ous bioac i e compounds, including phenolic
acids, la onoids, be alains, ca o enoids, and ocophe ols. These compounds ha e been
ound o ha e signi ican an ioxidan , -in lamma o y, -hype ensi e, and -hype glycaemic
and lipid-lowe ing p ope ies, and a e he main mechanis ic basis o hei bene icial e ec s
agains cance , neu odegene a ion, and me abolic diso de s such as obesi y and diabe es,
which a e majo isk ac o s o ca dio ascula disease [
375
]. By and la ge, quinoa and
ama an h a e gene ally conside ed sa e o consump ion and a e no commonly associ-
a ed wi h alle gies, al hough isola ed cases ha e been epo ed [
376
]. Howe e , mo e
esea ch is needed o ully unde s and he p e alence and mechanisms o ama an h and
quinoa alle gy.
Chia seeds also ha e excellen nu i ional p ope ies [
150
]. In addi ion o hei high
ib e con en and balanced ange o mac o- and mic o-nu ien s, chia seeds a e known o
hei high concen a ion o
α
-linolenic acid, which se es as a p ecu so o he syn hesis
o
ω
-3 a y acids wi h long chains such as DHA and EPA. I is now well es ablished ha
omega-3 polyunsa u a ed a y acids (PUFAs) con e se e al heal h bene i s. In pa icula ,
he consump ion o omega-3 a y acids has been associa ed wi h a dec ease in iglyce ide
and non-high-densi y lipop o ein choles e ol le els, suppo ing he p o ec i e ole o hese
Plan s 2024,13, 1914 25 o 45
a y acids agains ca dio ascula e en s [
377
]. In addi ion o educing dyslipidaemia,
omega-3 a y acids also exe p o ec i e bene i s agains a he oscle osis, in lamma ion,
diabe es, and cance , al hough some esea ch indings a e con o e sial and highligh he
need o mo e clinical s udies o cla i y he bene icial e ec s o
ω
-3 PUFAs [
378
]. Chia
seeds p o ide heal h bene i s due o bioac i e compounds such as phenolic molecules,
ocophe ols and ca o enoids. These compounds ha e an ioxidan p ope ies ha con ibu e
o heal h bene i s, pa icula ly in he igh agains diseases such as diabe es and cance [
224
].
Chia seeds also con ain phy os e ols, which a e mainly associa ed wi h educed choles e ol
and hepa op o ec i e e ec s. They also play a bene icial ole in diabe es and ca dio ascula
diseases [241].
In ecen yea s, he dis inc i e componen s and biological ac i i ies o mo inga ha e
been ex ensi ely s udied and documen ed, highligh ing i s po en ial heal h bene i s and
he apeu ic applica ions [
379
]. The whole M.olei e a plan , including lea es, ui s, pods,
lowe s, seeds, and oo s, exhibi s a ange o biological ac i i ies. Lea es a e ich in ac i e
cons i uen s such as la onoids, polyphenols, e penoids, phenylp opanoids, a y acids,
s e ols, and alkanes, as well as i amins and mine als [
380
–
383
]. Among hese, la onoids
and polyphenols a e he p ima y bioac i e molecules, showing an ioxidan , -cance , -
sepsis, -in lamma o y, -hype ensi e, and -diabe ic and hepa ic lipid-lowe ing p ope ies
and bene icial e ec s on obesi y- ela ed ep oduc i e diseases. In addi ion o a y acids,
essen ial amino acids, and la onoids, he seeds also con ain iso hiocyana es, which play
an impo an ole as an i-in lamma o y, -oxidan , -bac e ial, and -cance agen s [
384
].
Fu he mo e, newly disco e ed componen s ex ac ed om he s em o he plan , such as
icosanoic acid, choles -5-en-3-ol, s igmas e ol, and gamma-si os e ol, ha e shown po en
an i ungal ac i i y [
379
]. Po en an ioxidan s such as ocophe ols, asco bic acid, la onoids,
and ca o enoids ha e also been ex ac ed om M.olei e a lowe s [385].
Hemp seed, de i ed om he plan Cannabis sa i a L., has a ich his o y o use in Asia
da ing back o p ehis o ic imes. Recen ly, coun ies such as he Uni ed S a es, Canada,
and Aus alia ha e now legalized he a ming and use o hemp seed, p o ided ha i s
e ahyd ocannabinol con en emains below 0.3%. This legaliza ion has inc eased in e es
in hempseed due o i s ecognized nu i ional bene i s and pe cei ed pha maceu ical
applica ions [
386
,
387
]. His o ically, he economic impo ance o hemp has been in he
use o he ib e- ich s em, alued o p oduce ex iles, clo hing, and pape goods, while
i s seeds ha e emained la gely unde u ilized. Howe e , in ecen yea s he e has been a
su ge o in e es in explo ing he nu i ional and pha maceu ical p ope ies o hemp. In
pa icula , hempseed oil has he highes concen a ion o PUFAs o any ege able oil. I
is widely accep ed ha an inc eased in ake o PUFAs is associa ed wi h a educed isk o
se e al heal h condi ions, including ca dio ascula disease, cance , heuma oid a h i is,
hype ension, in lamma o y diso de s, and au oimmune diseases. O no e is he highly
desi able
ω
-6:
ω
-3 a io ound in hempseed, which ypically anges om 2.5 o 3.5/1, in
con as o he
ω
-6:
ω
-3 a io ound in ypical Wes e n die s, which no mally anges om 15
o 17/1. The impo ance o he
ω
-6/
ω
-3 a io in human heal h is well documen ed, wi h
a high a io (>10:1) in he human die being associa ed wi h an inc eased isk o cance ,
in lamma ion, and ca dio ascula disease. On he con a y, a a io lowe han 2.5:1 has been
associa ed wi h a educed isk o ch onic diseases, he supp ession o cance cells, and lowe
mo ali y [
184
]. In addi ion o i s oil con en , cannabis seeds con ain wo di e en phenolic
compounds: lignanamides and hyd oxycinnamic acids.
In i o
s udies ha e ex ensi ely
documen ed he an ioxidan p ope ies o hempseed lignanamides, including compounds
such as N- ans-ca eoyl y amine and cannabisin B. Addi ionally, hempseed is ich in
la onoids, which con ibu e signi ican ly o i s an ioxidan capabili ies, as demons a ed
by cell- ee sys em assays such as ORAC and FRAP. Fu he mo e, he di e en ac ions
ob ained om hempseed p ocessing show di e en le els o an ioxidan ac i i y, wi h he
coa ses ac ion, domina ed by hulls, showing he highes po ency in his ega d [
388
].
Hemp seed also con ains a highe concen a ion o ocophe ols han mos ege ables. These
ocophe ols ha e an ioxidan p ope ies ha e ec i ely inhibi he oxida ion o unsa u a ed
Plan s 2024,13, 1914 32 o 45
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Ja is, D.E.; Hoy, Y.S.; Ligh oo , D.J.; Schmöckel, S.M.; Li, B.; Bo m, T.J.A.; Ohyanagi, K.; Mine a, K.; Michell, C.T.; Sabe , N. The
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Pa i anage, D.S.R.; Rey, E.; Em ani, N.; Wellman, G.; Schmid, K.; Schmöckel, S.M.; Tes e , M. Genome-wide associa ion s udy in
quinoa e eals selec ion pa e n ypical o c ops wi h a sho b eeding his o y. eLi e 2022,11, e66873. [C ossRe ] [PubMed]
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Sandell, F.L.; Holzwebe , T.; S ee , N.R.; Dohm, J.C.; Himmelbaue , H. Genomic basis o seed colou in quinoa in e ed om
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andem duplica ions o SAD gene clus e s as d i e s o inc eased
α
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